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SPRUCE BUDWORM (Choristoneura Fumiferana)T PHEROMONE CHEMISTRY and BEHAVIORAL RESPONSES to PHEROMONE COMPONENTS and ANALOGS

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SPRUCE BUDWORM (Choristoneura Fumiferana)T PHEROMONE CHEMISTRY and BEHAVIORAL RESPONSES to PHEROMONE COMPONENTS and ANALOGS Journal of Chemical Ecology, Vol. 12, No. 2, 1986 SPRUCE BUDWORM (Choristoneura fumiferana)t PHEROMONE CHEMISTRY AND BEHAVIORAL RESPONSES TO PHEROMONE COMPONENTS AND ANALOGS PETER J. SILK and L.P.S. KUENEN Phe romone Research Group New Brunswick Research and Productivity Council P.O. Box 6000, College Hill Roarl Fredericton, New Brunswick E3B 5Hl (Received June 3, 1985; accepted August 1, 1985) Abstract-This paper reviews the sex pheromone chemistry and pheromone- mediated behavior of the spruce budworm and related coniferophagous (Choristoneura) budworms. ln C. funiferana, temporal changes in phero- mone-gland monounsaturated fatty acids (pheromone precursors) enable the prediction of the primary sex pheromone components. This technique may also be applicable for predicting additional pheromone components. Tetra- decanal (14:Ald), previously shown to enhance close-range precopulatory behavior, lowers the threshold of response by males for upwind flight to a pheromone-component source. Spruce budworm males maintain upwind flight to 95 : 5 (El Z)-1, 12-pentadecadiene (diolefi n analog) after initiating upwind flight to a primary-component pheromone source (95 : 5 ElZll-|4: Ald). 'lhis is the first demonstration of apparently normal male flight responses to a pheromone analog. Key Words-Cåoristoneura fumiferana, spruce budworm, Lepidoptera, Tortricidae, sex pheromone, behavior, flight tunnel, pheromone analog, pheromone fatty acid precursors. INTRODUCTION Lepidopteran female sex pheromones are, with few exceptions, multicompo- nent, consisting of a blend of two or more chemicals emitted at fairly consistent I Lepidoptera: Tortricidae 367 0098-033 I /86/0200 0367$05.00/0 O I 986 Plcnum Publishing Corporat¡on 368 Srlrc eNo KusNeÀr ratios and release rates. These chemical blends elicit in conspecific males se- quences of behaviors that include upwind flight which brings males to within close proximity of females. These behaviors are often readily observed in lab- oratory wind tunnels, and responses by males to synthetic pheromone sources can be compared to their responses to natural pheromone or females. The existence of a pheromone communication system in the spruce bud- worm (Choristoneura fumifurana Clemens) was demonstrated by Greenbank (1963), but elucidation of the blend of chemicals comprising the female sex pheromone is still incomplete. However, recent progress has been made in de- fining the sex-pheromone chemistry and pheromone-mediated behavior of this insect. This paper reviews the history and some current research related to the spruce budworm pheromone communication system. BACKGROUND In the genus Choristoneura (Lepidoptera: Tortricidae), known primary sex pheromone components are comprised of all-unsaturated c1a carbon-chain compounds. Different species' primary sex pheromone components have dif- ferent oxygenated functional groups and varying blends of these components (see Inscoe, 1982). Specifically, the coniferophagous spruce budworms com- prise a group of closely related (choristoneurø) species, native to North Amer- ica (Freeman,196l; Freeman and Stehr, 1967; powell, 1980). Six species have been studied in terms of their pheromone specificity (sanders, l97l; Sanders et al., 1914, 1977). The components of these female-produced sex pheromones form a group of congeneric aldehydes (Al1-14: Ald), acetates (Â11- l4:Ac), and alcohols (^11-14:OH),^11-Cr4 with blends, geometrical isomer ratios, and release rates specific to each species (Table l). In attempting to characterize the sex pheromone of budworm species through cross-attraction studies, Sanders et al. (1977) concluded that c. fumi- ferana, c. occidentalis, and c. bíennis apparently had similar pheromones while C. pinus pinus, C. orae, and C. retiniana (: C. viridis; powell, 1980) were mutually cross-stimulating but did not appear to have sex pheromone compo- nents in common with the former group. This was an accurate assessment; subsequent research has shown that the former group utilizes al l-14 :Ald's as primary sex pheromone components, whereas the latter, utilize Âll-14:Ac's or Â11-14 : Aclall-14:oH blends (Table l). It has yet to be verified that the c. biennis sex pheromone is a1l*14: Ald, although present evidence is strongly supportive (Sanders, 1971; Sanders et al., 1914, lgil). Sex pheromone com- ponents of the remaining five species have been documented: C. fumifurøna (Sanders and Weatherston, 1916; Silk et al., 1980) and C. occidentalis (Cory etal.,7982; Silk et al., 1982) have been shown to release Elzlt-\4:Ald as SPRTJCE BUDWORM PHEROMONE 369 TaeLp l. Spx Pnpnovo¡re CovtpoNeNtso op Cot¡I¡sRoPHACous Choristoneura spp. Species Primary Additional (Secondary) References C. fumiferana El l-14:Ald Weatherston et al., 197 | 96:4 E/Zll-| :Ald Sanders and Weatherston, 1976 95:5 E/Zll-14:Ald, l4:Ald Silk et al., 1980 Alford et at., 1983 C. occidentalis 92:8 E/Z|I-|4:Ald Cory et al., 1982 92:8 E/Zll-14:Ald 89:ll E/Z Silk et al., 1982 I I - 14:Ac Alford and Silk, 1983 85:15 E/Z 1l-14:OH C. biennis E/ZlI-14:Aldb Sanders, 1971 E/Z ratio unknown Sanders et al., 19'74 C. orae 82:9:9, Gray et al., 1984 Ell-14:Ac, Zll-14:Ac, El l-14:OH C. pinus pinus 90:10, Silk et al., 1985b 85:15 E/Zll-14:Ac, 85:15 E/Zll-14:OH C. retiniana 92:8, E or Zll-|4:OH Daterman et al., 1984 E/Zll-14:Ac (enhances trap capture) "E/Zll-\4:Ald : (E/Z)-ll-tetradecenal; E/Zll-l4:Ac : (E/Z)-11-tetradecenyl acetare. E/Z|1-14:OH : (E /Z) - 1 I -letradecen- I -ol ; I 4 : Ald : tetradecanal. áInferred from cross-attraction studies (Sanders, 197 l). primary sex pheromone components. In contrast, C. pinus pinus (Silk et al., 1985b), C. retiniana (Daterman et al., 1984), and C. orae (Gray et al., 1984) release EIZ ll-14: Ac or Elzll-|4: AclElZll-14:OH blends as primary com- ponents (Table l). However, all species have Â11-14 : Ac in common as the major component in the pheromone gland (in references above). In C. fumifurana All-tetrade- cenyl acetate is synthesized de novo only in the pheromone gland, the Al1- 14: Ac is the direct biosynthetic precursor to the aldehyde pheromone (Morse and Meighen , 1984). Morse and Meighen (1984) also found that the female diel aldehyde-emission period was synchronous with acetate production, and this supports their hypothesis that in this (coniferophagous) Choristoneura group a metabolic relationship exists between the aldehyde, acetate, and alcohol. It seems likely, therefore, that species speciflcity in pheromone production is con- trolled by species-specific metabolic processes, giving rise to different func- tional groups, ratios, and release rates from the common acetate precursor. 3',70 Slr-r eNn KUeNEN REVIEW OF SEX PHEROMONE CHEMISTRY OF SPRUCE BUDWORM Among coniferophagous budworms, the spruce budworm, C. fumifurana, has been the most intensively studied. Early work indicated that spruce bud- worm females release a pheromone that "attracts" males (Greenbank, 1963). Subsequently , Ell-14: Ald was identified (Wearherston et al. , 197 l) as a pher- omone component. The importance of adding the z to the E isomer was deter- mined upon reanalysis of female volatiles and a (96:4) E/Zll-14:Ald blend was shown to maximize trap captures (Sanders and weatherston, lg76). solvent extracts of excised pheromone glands were inactive in eliciting male response (Sanders, 1971). Reanalysis of gland extracts identified 811-14:OH (Weath- erston and Maclean, 1974) and Ell-14:Ac (Wiesner et al., 1979), both of which inhibited trap capture (Sanders and Lucuik, 1972; Sanders etal.,1972; Sanders 1976). More detailed chemical analyses (Silk et al., 1980) of pheromone gland extracts showed that A1l-14:Ac (20-40 ng/insecr), (l-3 ng/in- sect), and Al l-14: OH (1-3 ng/insecr) were all presenr^11-14:Ald in 95 :5 EIZ raÍios; the saturated analogs of each functionality were also present at ca. I % of the cor- responding E isomer. In contrast, effiuvia from "calling" females were found to contain Elzll-|4: Ald (95 :5; 10-40 nglinsect/nighr) and rhe saturated an- alog, tetradecanal (14:Ald) at ca.2% of the Z'll-14:Ald (Silk et at., l9g0). In addition, traces of (E)ll-14: Ac were found; no alcohols were detected. In the same study, field testing showed that there were no significant dif- ferences in trap captures between traps baited with all four components (for- mulated in PVC at female effiuvial ratios) compared to captures using the pri- mary components (95 :5 ElZll-l4:Ald) alone. The reduction of trap capture by admixture of All-14:Ac with 95:5 ElZll-|4:Ald (Sanders et al., 1972) was confirmed; however, this effect appeared to be negated by the presence of 14: Ald, but only when all components were present in synthetic sources in female effiuvial ratios (silk et at., 1980). Pheromone release rate by "calling" females, measured by a specific and sensitive bioluminescent assay technique (Morse eta1.,7982, Meighen et al., 1981, 1982), occurs mainly during sco- tophase in a series of "bursts" at rates as high as 50 ng/hr with considerable individual variability. REVIEW OF PHEROMONE-MEDIATED BEHAVIOR IN SPRUCE BUDV/ORM Behavioral pattems involved in mate location in feral insects generally in- volve upwind flight, apparent close-range orientation, and copulation (Roelofs and carde, 1977). As in most moths, spruce budworm males locate conspecific females by flying upwind along a pheromone plume. Often, the effects of pu- tative pheromone components on these male behaviors has been inferred from SPRUCE BUDWORM PHEROMONE 371 a chemical's effect on trap capture (until recently, this was also true for spruce budworm); however, observations and quantitation of some of these behaviors can be conducted in a sustained-flight wind tunnel (e.g., Miller and Roelofs, 1978). We review here recent field and wind-tunnel work, and present some recent progress.
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